Humpback whale song or humpback whale sonar? A reply to Au et al

Au et al.'s arguments against the hypothesis that humpback whale songs function as long-range sonar are based on questionable assumptions rather than on empirical data. Like other echolocating mammals (e.g., bats), singing humpback whales: 1) localize targets in the absence of visual information; 2) possess a highly innervated peripheral auditory system; and 3) modulate the temporal and spectral features of their sounds based on environmental conditions. The sonar equation is inadequate for determining whether humpback whale songs generate detectable echoes from other whales because it does not account for temporal variables that can strongly affect the detectability of echoes. In particular, the sonar equation ignores the fact that much of the noise encountered by singing humpback whales is spectrally and temporally predictable, and that audition in mammals is a dynamic and plastic process. Experiments are needed to test the hypothesis that singing humpback whales listen for and respond to echoes generated by their songs     

Combining data from a multisensor tag and passive sonar to determine the diving behavior of a sperm whale (Physeter macrocephalus)

Reports on the diving behavior of a sperm whale tagged and tracked on September 6, 2000 during the Sirena 2000 cruise in the Ligurian Sea. A total of about 4.5 h of acoustic and nonacoustic sensor data were recorded when a sperm whale was tagged with a Woods Hole Oceanographic Institution developed tag with a hydrophone, motion, and pressure sensors. The animal was simultaneously tracked with a passive sonar system deployed from the NATO research vessel NRV Alliance. By combining data from the tag and passive sonar, we were able to reconstruct a three-dimensional track of the whale, along with its orientation and vocal behavior. While it was tagged, the whale carried out three deep dives to a depth of about 900 m in an area with a bottom depth of about 2600 m. The inter-click intervals of the diving whale were not consistent with ranging on the bottom, but were consistent with the hypothesis that the whale was possibly echolocating on some target(s) near the depth at which it dove to feed. This study demonstrated an ability to track subtle changes in the behavior of diving whales. This ability is important for three areas: 1) basic research, 2) studies of the responses of these animals to controlled exposures of man-made noise, and 3) studies to infer the biological significance of behavioral disruption.     

Ecological aspects of fin whale and humpback whale distribution during summer in the Norwegian Sea

The Norwegian Sea is a migration and feeding ground for fin whales (Balaenoptera physalus) and humpback whales (Megaptera novaeangliae) in summer. During the last decade, significant structural changes in the prey community, including northerly expansion and movement in the distribution of pelagic fish species, have been reported from this ecosystem. However, little information on whale feeding ecology exists in the Norwegian Sea and surrounding waters. A total of 59 fin whales and 48 humpback whales were sighted during 864 h of observation over an observation distance of about 8200 nmi (15,200 km) in the Norwegian Sea from 15 July to 6 August 2006 and 2007. The fin whale group size, as mean (±SD), varied between one and five individuals (2.1 ± 1.2 ind.) and humpback whale group size varied between one and six individuals (2.5 ± 1.7 ind.). Fin‐ and humpback whales were observed mainly in the northern part of the study area, and were only found correlated with the presence of macro‐zooplankton in cold Arctic water. Humpback whales were not correlated with the occurrence of adult Norwegian spring‐spawning herring (Clupea harengus) except for the northernmost areas. Despite changes in the whale prey communities in the Norwegian Sea, no apparent changes in fin‐ or humpback whale distribution pattern could be found in our study compared to their observed summer distribution 10–15 years ago.     

Scale-dependent habitat use by a large free-ranging predator,the Mediterranean fin whale

Since the heterogeneity of oceanographic conditions drives abundance, distribution, and availability of prey, it is essential to understand how foraging predators interact with their dynamic environment at various spatial and temporal scales. We examined the spatio-temporal relationships between oceanographic features and abundance of fin whales (Balaenoptera physalus), the largest free-ranging predator in the Western Mediterranean Sea (WM), through two independent approaches. First, spatial modeling was used to estimate whale density, using waiting distance (the distance between detections) for fin whales along ferry routes across the WM, in relation to remotely sensed oceanographic parameters. At a large scale (basin and year), fin whales exhibited fidelity to the northern WM with a summer-aggregated and winter-dispersed pattern. At mesoscale (20–100 km), whales were found in colder, saltier (from an on-board system) and dynamic areas defined by steep altimetric and temperature gradients. Second, using an independent fin whale satellite tracking dataset, we showed that tracked whales were effectively preferentially located in favorable habitats, i.e. in areas of high predicted densities as identified by our previous model using oceanographic data contemporaneous to the tracking period. We suggest that the large-scale fidelity corresponds to temporally and spatially predictable habitat of whale favorite prey, the northern krill (Meganyctiphanes norvegica), while mesoscale relationships are likely to identify areas of high prey concentration and availability.     

Trends and interventions in large whale entanglement along the South African coast

The major causes of large whale entanglement in South Africa are static fishing gear, especially the type associated with the West Coast rock lobster Jasus lalandii industry, and large-mesh gillnets that are set off the coast of KwaZulu-Natal (KZN) to reduce shark attacks (shark nets). The prevalence of entanglements is seasonal with the peaks in activity coinciding with the breeding migrations of humpback whales Megaptera novaeangliae and southern right whales Eubalaena australis, the two large whale species that are the most prone to entanglement. Generalised linear models with a Poisson or quasi-Poisson distribution were used to describe the relationship between the number of incidents and time. Taking into account the combined length of shark-net installations per year as an offset variable, entanglement of humpback whales in shark nets increased at 15.1% per year (95% CI = 9.5–21.6%) from 1990 to 2009. This is comparable to the rate of increase in the numbers of this species migrating past the KZN coast, between 1988 and 2002 (9–11%). The number of reported incidents of southern right whales entangled in gear other than shark nets also increased between 1990 and 2009. This was accounted for by the increase in numbers of this species in South Africa (7% per year), so in neither case are the two species at increasing risk of individual entanglement, and anthropogenic factors including entanglement do not seem to be affecting the recovery of these whale populations. Nevertheless, there is concern regarding the vulnerability to entanglement of a small assemblage of humpback whales that habitually visits the West Coast in spring and summer. The continued recovery of whale populations is likely to lead to greater levels of anthropogenic interaction and heighten the need for adequate mitigation measures. The KZN Sharks Board and the South African Whale Disentanglement Network (since 2006) have respectively released (disentangled) 81% and 23% of confirmed entangled individuals, and recorded relevant information on entanglement incidents. Such information is critical for developing mitigation measures and monitoring the prevalence of entanglement.     

Acoustic interaction of humpback whales and whale-watching boats

The underwater acoustic noise of five representative whale-watching boats used in the waters of west Maui was measured in order to study the effects of boat noise on humpback whales. The first set of measurements were performed on 9 and 10 March, close to the peak of the whale season. The ambient noise was relatively high with the major contribution from many chorusing humpback whales. Measurements of boat sounds were contaminated by this high ambient background noise. A second set of measurements was performed on 28 and 29 April, towards the end of the humpback whale season. In both sets of measurements, two of the boats were inflatables with outboard engines, two were larger coastal boats with twin inboard diesel engines and the fifth was a small water plane area twin hull (SWATH) ship with inter-island cruise capabilities. The inflatable boats with outboard engines produced very complex sounds with many bands of tonal-like components. The boats with inboard engines produced less intense sounds with fewer tonal bands. One-third octave band measurements of ambient noise measured on 9 March indicated a maximum sound pressure level of about 123 dB re 1 microPa at 315 Hz. The maximum sound pressure level of 127 dB at 315 Hz was measured for the SWATH ship. One of the boats with outboard engines produced sounds between 2 and 4 kHz that were about 8-10 dB greater than the level of background humpback whale sounds at the peak of the whale season. We concluded that it is unlikely that the levels of sounds produced by the boats in our study would have any grave effects on the auditory system of humpback whales.     

Twelve years of tracking 52-Hz whale calls from a unique source in the North Pacific

A unique whale call with 50–52 Hz emphasis from a single source has been tracked over 12 years in the central and eastern North Pacific. These calls, referred to as 52-Hz calls, were monitored and analyzed from acoustic data recorded by hydrophones of the US Navy Sound Surveillance System (SOSUS) and other arrays. The calls were noticed first in 1989, and have been detected and tracked since 1992. No other calls with similar characteristics have been identified in the acoustic data from any hydrophone system in the North Pacific basin. Only one series of these 52-Hz calls has been recorded at a time, with no call overlap, suggesting that a single whale produced the calls. The calls were recorded from August to February with most in December and January. The species producing these calls is unknown. The tracks of the 52-Hz whale were different each year, and varied in length from 708 to 11,062 km with travel speeds ranging from 0.7 to 3.8 km/h. Tracks included (A) meandering over short ranges, (B) predominantly west-to-east movement, and (C) mostly north-to-south travel. These tracks consistently appeared to be unrelated to the presence or movement of other whale species (blue, fin and humpback) monitored year-round with the same hydrophones.     

Characterization of North Atlantic right-whale (Eubalaena glacialis) sounds in the Bay of Fundy

Hydrophone recordings, made in the presence of North Atlantic right whales (Eubalaena glacialis) in the Bay of Fundy during the daytime in July 1999, are used to determine the characteristics of the recorded sounds. A spectrogram-based method was implemented to discriminate whale sounds from background noise and the time-frequency envelope of the primary harmonic in the spectrogram was used as the basis for sound characterization. Sounds were typically (82%, n=45) in the 300- to 600-Hz range with up- and downsweeping modulations. Lower (<200 Hz) and higher (>900 Hz) frequency sounds were relatively rare. Most sounds were frequency modulated, with 95% of the observed instantaneous relative frequency variation being within /spl plusmn/4.5% of the mean peak frequency. Harmonics were observed in 18% of the sounds. The average sound duration was 0.42 s/spl plusmn/0.26 SD. The sounds occurred at a rate of between 0.3 and 0.7 min/sup -1/. The time intervals between adjacent sounds (2-700 s) were not randomly distributed. The number of sounds occurring among different waiting times did not reflect a Poisson distribution and a clustering of sounds (2 to 5 cluster/sup -1/) was observed. The sound characteristics are compared to those documented elsewhere and as reported for the southern right whale.     

Are solar activity and sperm whale Physeter macrocephalus strandings around the North Sea related?

In the final decades of the last century, an increasing number of strandings of male sperm whales (Physeter macrocephalus) around the North Sea led to an increase in public interest. Anthropogenic influences (such as contaminants or intensive sound disturbances) are supposed to be the main causes, but natural environmental effects may also explain the disorientation of the animals. We compared the documented sperm whale strandings in the period from 1712 to 2003 with solar activity, especially with sun spot number periodicity and found that 90% of 97 sperm whale stranding events around the North Sea took place when the smoothed sun spot period length was below the mean value of 11 years, while only 10% happened during periods of longer sun spot cycles. The relation becomes even more pronounced (94% to 6%, n = 70) if a smaller time window from November to March is used (which seems to be the main southward migration period of male sperm whales). Adequate chi-square tests of the data give a significance of 1% error probability that sperm whale strandings can depend on solar activity. As an alternative explanation, we suggest that variations of the earth's magnetic field, due to variable energy fluxes from the sun to the earth, may cause a temporary disorientation of migrating animals.     

Characteristics of the hepatic microsomal cytochrome P-450 system of the minke whale (Balaenoptera acutorostrata)

The whales are an interesting group in several aspects of science, including evolutionary biology, marine ecology and toxicology. Some of the whale species, including the minke whale, graze at the top of the food chains, where they become susceptible to the accumulation of marine pollutants ingested by organisms at lower levels. The northern stock of the minke whale grazes in Arctic waters in the summer, an area of increasing interest in the development of new oil fields. Studies on the metabolism, disposition and effects of xenobiotics in marine mammals are few. The only reports in the open literature so far have dealt with these questions in seals.^1–3 Although the marine mammals comprise a particularly interesting group in marine ecology and toxicology, no data have to our knowledge been presented on the xenobiotic metabolising enzyme systems of the whales. In this study, components of the microsomal cytochrome P-450 system were measured in liver samples from several females, one male, and two foetal minke whales, caught in the Norwegian whaling season of 1983. Because of the limited number of samples studied we have treated the samples individually and not performed any statistical analysis of the data. The trends discussed below must therefore be considered with this background.     

High gray whale relative abundances associated with an oceanographic front in the south-central Chukchi Sea

We describe gray whale (Eschrichtius robustus) distribution in the south-central Chukchi Sea in relation to environmental factors during two 5-day surveys in June and September of 2003. Whale counts per 10-min scan (an index of relative abundance) ranged from 0 to 41 in June and from 0 to 28 in September. CTD data showed an ocean front around 67.8°N with strong horizontal gradients in temperature, salinity, chlorophyll-a concentration and water-column stability. Highest whale abundance indices occurred in or near the front in both periods. Preliminary qualitative assessment of biological communities in the study area suggests that infaunal clams, echinoderms, euphausids, chaetognaths and Arctic cod were common, while ampeliscid amphipods, the previously abundant infauna (and likely prey) in the nearby Chirikov Basin feeding area, were not dominant. Euphausids may be a prey for gray whales in this area. We suggest that frontal systems may play an important role in eastern North Pacific gray whale foraging grounds. Further study is needed to fully describe the role of frontal systems in gray whale foraging grounds.     

Abundance of fin whales (Balaenoptera physalus) in the north-western Mediterranean Sea,using photo-identification and microsatellite genotyping

The Mediterranean fin whale population, Balaenoptera physalus, is resident, with almost no exchanges with the Atlantic population. The entire population was estimated at 1300 or 13,300 individuals by a recent project depending on the platform used. This disparity shows the importance of long-term monitoring with a unique protocol of survey. Capture-recapture approaches using dorsal photographs and genetic identity collections over a 10 years period were used to estimate the abundance of the north-western Mediterranean fin whale. We identified 332 individuals using photographs and 470 using genotypes, with a total of 546 individuals identified between 2008 and 2019, when some whales were double-marked. The inter-annual percentage of recapture varied between 15% and 17% respectively for genotypes and photographs methods. Using Cormack-Jolly-Seber models, the abundance of fin whales in the north-western Mediterranean is estimated at 1295 individuals (95% CI: 1116–1474) with a survival probability of 0.945 (95% CI: 0.690–0.993) from genotypes. Abundance estimates from combined collections (photographs and genotypes) and corrected photograph estimates were similar to the genetic ones. Future studies might prioritize the genetic approach which is the least biased and with a narrower confidence interval. The genetic abundance estimates show relative stability over time, when compared to 1990 estimates, and should be included in future conservation actions.     

Whale interactions with Alaskan sablefish and Pacific halibut fisheries: Surveying fishermen perception,changing fishing practices and mitigation

Whale depredation occurs when whales steal fish, damage fish or damage fishing gear. In Alaska, killer whales (Orcinus orca) and sperm whales (Physeter macrocephalus) primarily depredate on demersal sablefish (Anoplopoma fimbria) and Pacific halibut (Hippoglossus stenolepis) longline fisheries. Quantitative data on whale depredation in Alaska is limited due to low fishery observer coverage and minimal depredation evidence left on longline fishing gear. This study utilized semidirected interviews (n=70) and written questionnaires (n=95) with longline fishermen to examine: (1) perceptions and experiences of whale–fishery interactions in Alaska, (2) effects of depredation on fishing practices, and (3) potential depredation mitigation measures. Eighty-seven percent of fishermen surveyed agreed that whale depredation became worse between 1990 and 2010. Respondents reported changing their fishing practices in response to depredating whales in several ways, including: traveling up to 50 nautical miles and ceasing hauling operations up to 24 h until the whales left the fishing grounds. Respondents fishing in western Alaska, primarily encountering killer whales, were forced to wait longer and travel greater distances than fishermen operating in central and southeast Alaska, regions more affected by sperm whales. Deterrent research, gear modifications and real-time tracking of depredating whales were solutions favored by study participants. Survey respondent answers varied based on areas fished, quota owned, years involved in the fishery and vessel size. This study presents the first statewide evaluation of fishermen's perception and knowledge of whale interactions with the Alaskan longline fleet and is a critical step toward developing baseline data and feasible depredation mitigation strategies.     

Against the humpback whale sonar hypothesis

A rebuttal is presented to the article of L.N. Frazer and E. Mercado (ibid., vol. 25, pp. 160-182, 2000), who presented a sonar model for humpback whale song, is presented. This rebuttal considers the noise-limited form of the sonar equation, current understanding of humpback whale behavior and the characteristics of humpback whale songs, along with arguments from an evolutionary perspective. Arguments from all of these different aspects do not support the model of Frazer and Mercado     

Qualitative correlation of marine mammals with physical and biological parameters in the Ligurian Sea

In support of its acoustic risk mitigation policy, NATO SACLANT Undersea Research Centre (SACLANTCEN) is sponsoring a series of sea trials, entitled "Sirena" to collect a multiyear integrated oceanographic, biological, and hydrographic data set, the goal being to explain, based on these parameters, the distribution of marine mammals found in specific locations. By understanding how ocean dynamics affects the distribution and behavior of whales and the organisms forming the food web upon which the whales feed, it may be possible to conduct acoustic exercises in areas of low cetacean density. The first two Sirena multidisciplinary cruises were conducted in the Ligurian Sea in late summer time frame during 1999 and 2000. The focus of this analysis is to determine whether remotely sensed satellite data can indicate nutrient-rich regions in areas where the oceanography is known and to determine if these regions of higher productivity, coupled with knowledge of cetacean presence from all available sources, could be used as an indicator of marine mammal presence for acoustic risk mitigation purposes. For the two years of data examined, cooler sea-surface temperature data correlated with high levels of chlorophyll production as seen by remotely sensed images. This remotely sensed data correlated well with measured subsurface values of the same parameters. Coincident sightings of three species of marine mammals indicated that fin and sperm whales generally preferred the deep, nutrient-rich portion of the basin while Cuvier's beaked whales preferred a submarine canyon where there is a frontal influence, as indicated from satellite data and historical oceanography. This paper is intended as a contribution to the longer term objective of developing the means to accurately predict cetacean presence from physical oceanographic characteristics.     

Intra- and inter-species differences in persistent organic contaminants in the blubber of blue whales and humpback whales from the Gulf of St. Lawrence, Canada

Biopsy samples of blubber from adult male and female blue whales, and from female and young-of-the-year humpback whales were collected during the summers of 1992-1999 in the Gulf of St. Lawrence, Canada. In blue whales, concentrations of 25 PCB congeners, DDT and metabolites and several other organochlorine compounds were present at higher concentrations in the blubber of males relative to females; reflecting maternal transfer of these persistent contaminants from females into young. Sex-related differences in concentrations were not observed with less persistent contaminants, such as HCHs. In humpback whale samples, there were no significant differences in the concentrations of PCBs and organochlorine compounds in the blubber of females and calves. These data indicate that calves quickly bioaccumulate contaminants by transplacental and lactational routes to concentrations that are in equilibrium with females. In comparisons between contaminant concentrations and patterns in the blubber of female blue and humpback whales, there were no significant differences in concentrations, but the proportions of some PCB congeners, HCH isomers, and DDT and its metabolites were different in the two baleen whale species. These may reflect differences in the diet of the two species, since fish comprise a large part of the diet of humpback whales and blue whales feed exclusively on euphausiid crustaceans (i.e. krill).     

Acoustic behaviour of southern right whales in relation to numbers of whales present in Walker Bay,South Africa

Estimating numbers of whales present in an area from recorded call rates could be a useful conservation tool. We recorded southern right whale Eubalaena australis vocalisations and presence in Walker Bay on the south coast of South Africa. In all, 45 sessions with synchronous acoustic and visual data were analysed to determine call rates directly (overall call rate, OCR) and in relation to the number of whales sighted (call rate per whale, CPW) and number of groups sighted (call rate per group, CPG). The OCRs were examined in the presence of varying numbers of whales, using a log-linear model to investigate the dependence of the call rate on whale density. The number of whales present exerted a strong quadratic effect on the OCR, which peaked at around 15 whales and decreased to a low rate as whale presence approached a maximum, for all calls combined and for four of their constituent 13 call types: a quadratic trend was present to varying degrees among the remaining call types. Both quadratic and linear trends were absent when OCR was assessed against number of groups present, possibly because group size increased with increasing density of whales. A linear regression on the CPW and CPG suggested that there was a negative inverse relationship with the number of whales and groups present respectively. These findings are important in that, while they preclude the estimation of absolute numbers from call rates, they imply that, under the conditions prevailing in Walker Bay, southern right whales were aware of the presence, arrival and departure of other animals in their vicinity and adjusted their vocal behaviour accordingly.     

Distribution and seasonal abundance of large cetaceans in the Durban whaling grounds off KwaZulu-Natal,South Africa, 1972–1975

Daily charts of the aerial search effort (432 206 nautical miles) of the Union Whaling Company and 1 099 sightings of 10 497 whales were available from 628 flights off Durban between 1972 and 1975. Densities of whales were analysed by month and water depth distribution over the four-year period. Low observed densities of blue Balaenoptera musculus, right Eubalaena australis, sei B. borealis and humpback Megaptera novaeangliae whales most likely resulted from earlier whaling pressure. Seasonality of blue, sei and humpback whales was bimodal, indicative of winter migrations to the north of the Durban whaling grounds, whereas the unimodal seasonality of fin whales B. physalus and minke whales B. bonaerensis or B. acutorostrata suggest the offshore region as the northern terminus of their migrations. Sperm whales Physeter macrocephalus migrate northwards offshore of the KwaZulu-Natal coast in autumn/early winter and southwards in late winter/spring, with larger males migrating later than the smaller males and females. Killer whale Orcinus orca presence was coincident with that of offshore minke whales and the southward migrations of other baleen whales, whereas densities of animals deemed as bottlenose whale Hyperoodon planifrons suggest strong early and late summer seasonal abundance in the offshore region. Such extensive surveys offshore of the KwaZulu-Natal coast are unlikely to be repeated; hence, data-extraction of whaling records provides a valuable source of seasonal and distributional information for marine management.     

Long‐term presence and habitat use of Cuvier's beaked whale (Ziphius cavirostris) in the Central Tyrrhenian Sea

Cuvier's beaked whale (Ziphius cavirostris, G. Cuvier 1823) is a poorly known species and many international agreements have asked for a better understanding of its biology for conservation purposes. In the present study, systematic cetacean surveys were carried out from ferries along a trans‐regional fixed transect in the Central Tyrrhenian Sea (Civitavecchia, Latium – Golfo Aranci, Sardinia), just outside the southeastern border of the Pelagos Sanctuary. This research provided long‐term, consistent data on Cuvier's beaked whale during two research periods (1990–1992 and 2007–2011). The objective of the research was to compare the presence, distribution and habitat use of Cuvier's beaked whale between the two investigated periods. Summer data (June–September) from the two periods were compared in terms of frequency of sightings, group size and spatial distribution related to the main ecogeographical features. A presence‐absence model (generalized additive modelling) was performed to predict habitat suitability in the two study periods. The results highlight long‐term site fidelity of Cuvier's beaked whale in the Central Tyrrhenian Sea with encounter rates comparable to the ones reported for other key areas. Separate suitability models based on 1990s and 2000s data appeared to work for each individual time period but differences were evident between the two periods, indicating changes in habitat selection over time. Our findings of the study appear to expand the definition of suitable beaked whale habitat and underline how the temporal scale of the analysis can affect the results in habitat studies. Moreover, this research highlights the importance of the Central Tyrrhenian Sea marine region for Cuvier's beaked whale and the ability of continuous monitoring to identify changes in cetacean frequency and distribution, necessary for adaptive conservation management approaches.